vol. 172, no.
6 the american naturalist december 2008
Disintegration of the Ecological Community
American Society of Naturalists Sewall Wright Award Winner Address *
Robert E. Ricklefs†
Department of Biology, University of Missouri, St. Louis, Missouri
63121
abstract: In this essay, I argue that the seemingly indestructible
concept of the community as a local, interacting assemblage of species
has hindered progress toward understanding species richness at local
to regional scales. I suggest that the distributions of species within
a region reveal more about the processes that generate diversity pat-
terns than does the co-occurrence of species at any given point. The
local community is an epiphenomenon that has relatively little ex-
planatory power in ecology and evolutionary biology. Local coexis-
tence cannot provide insight into the ecogeographic distributions of
species within a region, from which local assemblages of species
derive, nor can local communities be used to test hypotheses con-
cerning the origin, maintenance, and regulation of species richness,
either locally or regionally. Ecologists are moving toward a com-
munity concept based on interactions between populations over a
continuum of spatial and temporal scales within entire regions, in-
cluding the population and evolutionary processes that produce new
species.
Keywords: biodiversity, biogeography, community ecology.
Recent symposia of the Ecological Society of America
(Webb et al. 2006), the American Society of Naturalists
(Harrison and Cornell 2007), and the British Ecological
Society (Speciation and Ecology, Sheffield, March 29–30,
2007) have been devoted to phylogenetic and geographic
approaches to the study of ecological systems, including
ecological communities. This broadening perspective,
* Robert E. Ricklefs received the 2005 Sewall Wright Award. The Sewall Wright
Award honors a senior but still active investigator who is making fundamental
contributions to the Society’s goals, namely, promoting the conceptual uni-
fication of the biological sciences.
†
E-mail: ricklefs@umsl.edu.
Am. Nat. 2008. Vol. 172, pp. 741–750. 䉷 2008 by The University of Chicago.
0003-0147/2008/17206-50514$15.00. All rights reserved.
DOI: 10.1086/593002
linked as it is to such disparate issues as global climate
change and molecular phylogenetics, has stimulated ecol-
ogists to consider with more interest the history of the
environment and the historical and geographic contexts
of ecological systems (Latham and Ricklefs 1993a; Wiens
and Donoghue 2004; Jaramillo et al. 2006; Ricklefs et al.
2006). We appear to be in the midst of a major synthesis
in ecology (Lawton 1999), comparable to the maturation
of ecosystem perspectives during the 1950s (McIntosh
1985) and population perspectives during the 1960s (Mac-
Arthur 1972; Kingsland 1985).
Despite these developments, however, ecologists, for the
most part, continue to regard local communities as eco-
logical units with individual integrity (Harrison and Cor-
nell 2008). Empirical and experimental studies, including
recent analyses of food webs and mutualistic networks
(Jordano et al. 2003; Lewinsohn et al. 2006), circumscribe
populations and communities locally (Morin 1999; Chase
and Leibold 2003). Spatial scale rarely appeared in “com-
munity” theory until recently (Ives and May 1985; Brown
et al. 2000; Leibold et al. 2004; McCann et al. 2005), and
where it does appear, it is generally limited to the influence
of dispersal limitation and population aggregation on local
coexistence (Belyea and Lancaster 1999; Chesson 2000).
The recent review by Agrawal et al. (2007) on “filling key
gaps in population and community ecology” recognized
some implications of the spatial and historical contexts of
local “communities” (see Thompson et al. 2001) but em-
phasized the central role of species interactions on local
scales in limiting coexistence. This is particularly signifi-
cant because the Agrawal et al. review, with 16 prominent
ecologists as authors, was commissioned by the National
Science Foundation to recommend research priority areas
in population and community ecology.
Evidence versus Belief in Community Ecology
In spite of decades of evidence to the contrary, ecologists
have been reluctant to abandon a local concept of the
742 The American Naturalist
community (Ricklefs 1987; Harrison and Cornell 2008).
It has been eight decades since Gleason (1926) challenged
Clements’s (1916) perception of the community as an in-
tegral unit in ecology, and more than 50 years have passed
since Whittaker’s (1953, 1967) definitive work on the dis-
tributions of plant species across ecological gradients. In
spite of the influence of MacArthur and Wilson’s (1963,
1967) theory of island biogeography, in which colonization
of islands from external sources was a primary driver of
diversity, ecologists broadly began to accept the influence
of regional processes on the species membership of local
ecological assemblages only about 25 years ago (Hanski
1982; Cornell 1985; Ricklefs 1987; Pulliam 1988; Stevens
1989; Kareiva 1990; Cornell and Lawton 1992; Hanski and
Gilpin 1997; Srivastava 1999). During this time, meta-
population ecology (Hanski and Gilpin 1997), landscape
ecology (Turner et al. 2001), the mosaic theory of coevo-
lution (Thompson 2005), neutral theory (Hubbell 2001;
Hubbell and Lake 2003), macroecology (Brown 1995; Gas-
ton and Blackburn 2000; Blackburn and Gaston 2001),
and metacommunity perspectives (Holyoak et al. 2005),
among other developments, have embraced large-scale
patterns and processes.
New ideas and perspectives sometimes penetrate a dis-
cipline slowly, but ecology seems to have been especially
resistant to the disintegration of the community as a cen-
tral concept. (I use “disintegrate” in the sense of breaking
an entity into parts or components or dispersing its co-
hesion or unity.) My own preoccupation with this idea
might be misplaced, but Lawton (1999, p. 183) also em-
phasized, somewhat pessimistically, that “the major weak-
ness of traditional community ecology, and why it has so
conspicuously failed to come up with many patterns, rules,
and workable continent theory, is its overwhelming em-
phasis on localness.” Perhaps strong local interactions are
sufficient for an understanding of the structure of species
assemblages, giving local communities, as well as the local
populations they comprise, a primary ecological validity
(MacArthur 1965, 1972; Cody 1974; Huston 1994; Belyea
and Lancaster 1999; Weiher and Keddy 1999). Nonethe-
less, the persistence of the idea of local community integ-
rity brings to mind two additional possibilities. One is a
limitation of language. The words “population” and “com-
munity” have generally accepted definitions that corre-
spond to entities, just like the words “dog” and “house,”
neither of which would withstand disintegration and still
retain its essential features. The second is a problem of
utility. Disintegrating “community” might be justified, but
would this inform our science? That is, would significant
and compelling new theory and research programs ma-
terialize? I argue here that local coexistence can be un-
derstood only in terms of the distributions of species
within entire regions, which are determined by diversifi-
cation and adaptation within the regional ecological space
in combination with the interactions of species over entire
regions (Ricklefs 2007b).
Local Communities Are Not Integral Entities
Ecologists consistently define communities as units: gen-
erally, populations of different species living within a spec-
ified location in space and time. Although this definition
need not connote an integral nature, common usage in
ecology implies (artificially) bounded units that may be
treated as entities. These units are also fixed in a hierarchy
of organism-population-community-ecosystem-biosphere
(Lidicker 2008) and are generally taught in this order in
introductory ecology courses (e.g., Molles 2005; Smith and
Smith 2006; Cain et al. 2008). The idea that a local as-
semblage consists of species with partially overlapping dis-
tributions that happen to co-occur at a point—Gleason’s
(1926) individualistic concept of the community—does
not easily fit into this hierarchical concept, for example,
in the way that species fit into genera and genera make
up families. Ecologists generally accept that populations
have a spatial extent and are integrated by the movement
of individuals within them (Pulliam 1988, 2000; Loreau
and Mouquet 1999; Amarasekare and Possingham 2001).
From this perspective, a local community cannot be in-
clusive of the populations of its component species.
Of course, populations have geographic structure de-
fined by barriers to dispersal, which result in interrupted
gene flow, isolation by distance, and local genetic and eco-
typic differentiation. Sewell Wright (1978) made seminal
contributions to understanding spatial population struc-
ture. Dispersal limitation has infused work on species in-
teractions as well, for example, through Thompson’s geo-
graphic mosaic of coevolution (Fox and Morrow 1981;
Thompson 2005; Siepielski and Benkman 2007) and Hub-
bell’s metacommunity construct (Hubbell 2001; Condit et
al. 2002; Chave 2004; Holyoak et al. 2005), which parti-
tions ecological systems into local and global entities con-
nected by a migration parameter (m). Dispersal limitation
is necessary for allopatric speciation (Coyne and Orr 2004;
Price 2008), a process that is often linked to global vari-
ation in species richness (Mittelbach et al. 2007). Dispersal
limitation within metacommunities produces a theory of
species-area relationships and beta diversity (Hubbell
2001; Condit et al. 2002). Regardless of the degree to which
populations are subdivided, however, they exhibit inte-
gration over spatial scales that greatly exceed the generally
accepted extent of local communities.
Disintegration of local communities, made necessary by
the spatial extent of their component populations, frees
one to address the geographic distributions of populations
and the historical development of local assemblages of

species (Ricklefs 1989; Webb et al. 2002; Cavender-Bares
et al. 2006). One can then ask whether assemblages achieve
local equilibria or, conversely, whether local diversity re-
flects regional rates of species production and extinction
(Cardillo 1999; Cardillo et al. 2005; Allen et al. 2006; Rick-
lefs 2006b), constraints on adaptation within ecological
space (Wiens and Donoghue 2004; Ricklefs 2006a), and
dispersal limitation through geographic space (Svenning
and Skov 2004; Vormisto et al. 2004). Regional perspec-
tives reveal departures from local equilibria, discussed be-
low, that bring into question the development and testing
of theory based on local coexistence.
Disintegration raises the issue of scale in ecology (Allen
and Starr 1982). When boundaries break down and pattern
and process lack discrete spatial extent, time and space
become continuous (Shurin and Srivastava 2005). Within
large regions—continents or major ocean basins—pro-
cesses with different extents in time and space influence
the diversity of species over a range of scales. At the “re-
gional” end of the spectrum, matching the distributions
of entire species, allopatric speciation is the creative engine
of species richness (Cardillo et al. 2005; Jablonski et al.
2006; Ricklefs 2006b; Roy and Goldberg 2007; Weir and
Schluter 2007); shifts in topography and climate, as well
as catastrophic events with regional effects, can also reduce
species richness at these large spatial scales. At the “local”
end of the spectrum, matching the dispersal distances of
individuals within populations, competitive and other in-
teractions between species assume greater importance
(MacArthur 1970; Brown 1981; Brown et al. 2000). In-
tervals between speciation and catastrophic events, as well
as the intervals required for substantial climatic and to-
pographic change, are much longer than the intervals be-
tween births and deaths that determine local changes in
population size (Schneider 1994, 2001).
History, Geography, and the Community Concept
MacArthur (1965) argued that one could ignore large-scale
history and geography in the study of ecological com-
munities because local processes influencing the coexis-
tence of species (i.e., competition, predation, and mutu-
alism) come into equilibrium locally so rapidly that
processes on larger scales are inconsequential. Thus, local
assemblages have limited membership determined by spe-
cies interactions (MacArthur and Levins 1967; Vandermeer
1972). Accordingly, variation in diversity should reflect the
way in which physical conditions influence the coexistence
of species, leading to the prediction that species richness
should vary in direct relationship to the environment, par-
ticularly climate (Ricklefs 1977; Currie 1991; O’Brien 1998;
Hawkins et al. 2003; Currie et al. 2004; Kilpatrick et al.
2006). A second important prediction from this idea is
The Ecological Community 743
that local species richness should be independent of re-
gional species richness (Terborgh and Faaborg 1980), in
which case discrepancies between the two would be ex-
plained by differences in species turnover among habitats
or over distance: beta diversity (Cody 1975).
The recent literature in ecology has addressed these pre-
dictions in detail, producing two general conclusions. First,
species richness is generally related to physical conditions
of the environment—temperature and water availability,
for example (Hawkins et al. 2003)—although diversity on
both regional and local scales can differ between regions
with similar environments (Ricklefs and Latham 1993; Hu-
gueny et al. 1997; Qian and Ricklefs 2000; Ricklefs et al.
2006). The diversity-environment correlation need not im-
ply local equilibrium with respect to physical conditions.
A close relationship between the two is also predicted by
hypotheses based on evolutionary diversification out of
ecological zones of origin (Terborgh 1973; Latham and
Ricklefs 1993b; Wiens and Donoghue 2004; Ricklefs
2006a). Second, where tests are feasible, generally in in-
tercontinental comparisons, local species richness is di-
rectly related to regional species richness (Fjeldså and Lov-
ett 1997; Srivastava 1999; Ricklefs 2000; Rahbek and
Graves 2001; Shurin and Srivastava 2005; Rahbek et al.
2007; see Hillebrand and Blenckner 2002 concerning the
value of these tests). This suggests that large-scale processes
(species production and regional extinction) influence
both regional and local diversity or, alternatively, that en-
vironmental conditions similarly influence processes on
both regional and local scales. These alternatives can be
tested statistically with structural equation modeling (e.g.,
Harrison et al. 2006). Differences in diversity in similar
environments in different regions—so-called diversity
anomalies—imply that large-scale regional and historical
factors can influence local species richness over and above
the influence of the local environment (Latham and Rick-
lefs 1993a; Ricklefs and Latham 1993; Qian and Ricklefs
2000; Ricklefs et al. 2006).
According to the theory of ecological drift (Hubbell
2001), the equilibrium number of species within a meta-
community is a function of the speciation rate and the
total number of individuals, which generally varies with
the size of the region. If speciation rate were influenced
by environmental conditions (Davies et al. 2004; Allen et
al. 2006), this theory also would produce diversity-envi-
ronment relationships, as well as a correlation between
diversity and the size of a region (Terborgh 1973; Rosen-
zweig 1995; Ricklefs 2006b). Hubbell’s model also predicts
a direct relationship between the number of species in a
local assemblage and regional diversity, determined by the
balance between species production and dispersal. Eco-
logical drift ignores obvious ecological factors—particu-
larly environmental variation and habitat specialization—
744 The American Naturalist
but nonetheless emphasizes the connection of local di-
versity to large-scale processes. Alas, community drift is
much too slow to account for observed rates of species
turnover (Leigh 1981, 2007; Nee 2005; Ricklefs 2006c),
just as genetic drift is slow compared to adaptive evolution.
Consequently, only more powerful evolutionary and eco-
logical forces can account for patterns of species richness.
The Regional Community Concept
How can we reconcile these observations: diversity-envi-
ronment relationships, local-regional diversity relation-
ships, and regional effects (diversity anomalies)? By re-
placing artificial local-community boundaries with a
time-space continuum of process and pattern in popula-
tion and community ecology (Shurin and Srivastava 2005),
we might imagine processes of large and small extent tend-
ing toward equilibrium over a continuum of scale. Ac-
cordingly, distinguishing within-habitat and between-hab-
itat components of diversity would become arbitrary and
lose meaning. This applies to Whittaker’s (1960, 1972)
alpha diversity, which addresses the number of species
within a particular area, community, or ecosystem. Alpha
diversity may be defined for an area as small as a local
plot or as large as an entire region, at which point it is
identical to regional, or gamma, diversity. Beta diversity,
which describes the rate of change, or turnover, in species
composition across habitats or among communities, thus
loses meaning because it depends on the scale of the alpha
component of diversity, emphasizing the continuity of pat-
tern over space (Loreau 2000) or time.
The “open-community” perspective of Gleason (1926),
supported by Whittaker’s depictions of the distribution of
populations over ecological gradients within regions
(Whittaker 1967), signaled a major shift of emphasis that
has yet to materialize fully, in spite of the widespread
application of ordination and other approaches to han-
dling spatial variation in ecology (Legendre and Legendre
1998). The local community concept represents what I
think of as a vertical perspective on species occurring
within an arbitrarily bounded area, vertical because the
species present locally add together—pile up, so to speak—
to form the local assemblage. The occurrence of species
elsewhere within a region is of no consequence. In con-
trast, the regional community concept corresponds to a
horizontal perspective on the distribution of populations
over ecological and geographic gradients (e.g., Pulliam
2000; Pyke et al. 2001; Phillips et al. 2003; Wiens et al.
2007; Butt et al. 2008). Each distribution reflects the eco-
logical heterogeneity of a region, the adaptations of in-
dividuals to physical conditions, vegetation structure, and
so on (i.e., the fundamental niche), the ability of individ-
uals to disperse to isolated suitable areas, and the pro-
ductivity of a population as it is influenced by its evolved
interactions with resources, competitors, and consumers
(i.e., the realized niche).
Developing a Regional Perspective
From the distributions of species along environmental or
geographic gradients, the local community can be reduced
to a single point shared by many species. Lacking spatial
extent, this “point community” ceases to be an entity. The
community concept is replaced by the spatial distributions
of populations, which now become the primary focus for
understanding biodiversity patterns. Accordingly, the beta
(spatial) component of diversity is defined by the extent
of each population over space or over an ecological gra-
dient, shifting our focus from the local community to the
individual population within the region.
Imagine an ecological gradient of length V harboring
species (populations) that occupy average length v. The
probability that a particular species occurs at a particu-
lar point is v/V, and the average point diversity is thus
S # (v/V ), where S is the number of species in the region.
Point diversity varies in direct relation to both S, reflecting
large-scale processes that determine regional species rich-
ness, and v, the outcome of processes that determine the
ability of a population to spread and maintain itself over
an ecological or spatial gradient. These processes include
species interactions over the entire region. One outcome
of these interactions is that each population will approach
a stable “carrying capacity” over the region as a whole,
being confined by consumers and competitors to favorable
parts of the ecological space in which births plus immi-
gration balance deaths plus emigration, on average. The
resulting demographic equivalence of all species within a
region allows slow processes, such as the production of
species, to influence S; to the extent that higher S results
through regional competition in smaller v, regional pro-
cesses influence point diversity as well (Ricklefs 1989).
To adopt a regional, or horizontal, perspective, ecolo-
gists must turn their attention away from local assemblages
and toward populations, which, while not completely dis-
crete entities, interact with other populations over their
entire distributions. Ecologists have yet to develop a theory
tying together the ecogeographical distribution and local
abundance of populations, although this is a key issue in
the agenda of macroecology (Brown 1995; Gaston and
Blackburn 2000; Gaston 2003) and, to some extent, meta-
community ecology (Holyoak et al. 2005). At the core of
this issue is the concept of the niche, referring to the
occupation of ecological space (Brown 1981, 1984; Pulliam
2000; Chase and Leibold 2003). In locally defined com-
munity ecology (the vertical perspective), the niche reflects
evolved attributes of individuals for exploiting resources.

The horizontal perspective implies an additional, between-
habitat, or population, component to the niche, shaped
by the spatial dynamics of populations that link interac-
tions among species over large regions.
Filling Regional Ecological Space
Can the regional-horizontal perspective produce new in-
sights and promote the development of new theory in
ecology? A pessimistic answer would be that spatial com-
ponents of population processes and consideration of both
individual and population components of niche space cre-
ate unmanageable complexity (Lawton 1999; McPeek and
Brown 2007). The dynamics of species production and
evolutionary adaptation, as well as the causes and uncer-
tainties of extinction, pose additional challenges. Diversity
unfolds within a region as evolving clades of species grow
and shrink. Evolutionary conservatism of ecological re-
lationships (Westoby et al. 1995; Ackerly 2003, 2004; Wes-
toby 2006) constrains the way that species fill regional
niche space (Webb et al. 2002; Cavender-Bares et al. 2004;
Silvertown et al. 2006a, 2006b).
An important issue is whether, barring environmental
catastrophe, the number of species within a region typically
achieves equilibrium, with balanced rates of speciation and
extinction. Is diversity stably regulated within regions?
Analyses of changes in diversification rates based on phy-
logenetic data (Pybus and Harvey 2000; Nee 2006; Rabosky
et al. 2007) suggest that many clades show early bursts of
diversification followed by relative quiescence (e.g., Lov-
ette and Bermingham 1999; Harmon et al. 2003; Kozak
et al. 2006).
Analyses of diversity in the fossil records of several
groups have revealed relative constancy in the number of
species over periods of tens of millions of years (e.g., Alroy
2000; Jaramillo et al. 2006; Alroy et al. 2008). The inde-
pendence of clade size and age reported in flowering plants
(Ricklefs and Renner 1994; Magallón and Sanderson
2001), birds (Ricklefs 2006b), and squamate reptiles (Rick-
lefs et al. 2007; cf. McPeek and Brown 2007) further in-
dicates that the regional diversity within individual clades,
and perhaps within entire biotas, is regulated, at least rel-
ative to the turnover of species within clades (Ricklefs
2007a, 2009; Rabosky and Lovette 2008). Changing en-
vironmental conditions or the appearance of key evolu-
tionary innovations might result in the ascendancy of a
particular clade at the expense of others (Rabosky et al.
2007). Such trends are slow compared to intervals between
lineage splitting and extinction but much more rapid than
predicted from the dynamics of random events (Ricklefs
2007b).
The Ecological Community 745
The Regulation of Species Richness
The regional perspective challenges us to understand the
factors that determine the number of species within a re-
gion and how this number is linked to the geographic and
ecological distributions of individual species. The absence
of a diversity-age relationship among clades suggests that
regions have carrying capacities for species as well as for
individuals. Carrying capacities for populations can be
linked to the relationship between resource availability and
individual resource requirement. A carrying capacity for
number of species is more problematic. Ricklefs (2009)
found that the numbers of species in orders of flowering
plants and tribe- to family-level clades of passerine birds
were correlated between continents. This result suggests
that, to the extent that diversification of these clades has
been independent in the different regions, each clade
achieves a similar equilibrium number of species. Ac-
cordingly, not only would the total number of species
within a region be regulated but individual clades also
would appear to be constrained within the total regional
ecological space.
The size of a clade tends to vary in proportion to the
size of the region in which it has diversified (Ricklefs
2006b; Ricklefs et al. 2007). If clade size were a regional
property, it could be determined by increased rates of spe-
cies production (and less frequent extinction) in a larger
regional area or by the ability of large regions to accom-
modate more species when physiographic and ecological
complexity provides opportunities for allopatry and pop-
ulation turnover. Equilibrium island biogeography theory
(MacArthur and Wilson 1967) states that diversity re-
sponds to the pressure of colonization and also reflects
island-size–dependent extinction (e.g., Ricklefs and Ber-
mingham 2002, 2004). When islands are large enough,
internal species production further elevates diversity in-
dependently of an island’s range of environments (Losos
and Schluter 2000). Thus, in island systems, both local
and regional (pisland) diversity reflect forces that are ex-
ternal to the ecological characteristics of local habitats or
the habitat heterogeneity of an island. We need to extend
our spatial scale only slightly to suppose that continental
biotas come under the influence of similar considerations.
Ecologists understand in detail the foundation mecha-
nisms of evolutionary adaptation, physiological limits to
distribution and production, population dynamics and
species interactions, and the trophic structure of ecosys-
tems. However, the expression of these mechanisms in a
complex world with a long history is bound to be idio-
syncratic and unpredictable, just as evolutionists, with
their detailed understanding of the mechanisms of heredity
and selection, might fail to predict the existence of kan-
garoos, for example. Nonetheless, nature, including its di-
746 The American Naturalist
versity, does exhibit pattern, and it should be possible to
understand the origin of that pattern.
The Future of the Community Concept
I return to the question posed earlier in this essay, namely,
can the disintegration of the concept of “community” lead
to new theory and research? My answer echoes E. O. Wil-
son’s (1959, p. 122) admonition a half-century ago that
there is “a need for a ‘biogeography of the species,’ oriented
with respect to the broad background of biogeographic
theory but drawn at the species level and correlated with
studies on ecology, speciation, and genetics.” The local
“community” consists of those species whose distributions
include a particular point in space and time. However, the
integral units of community organization are the popu-
lations of species within regions that might or might not
encompass that point. Indeed, the presence and relative
abundance of a species at a particular point might depend
on interactions with populations that do not occur there.
Thus, to understand the coexistence of species locally, one
must understand what shapes species distributions within
regions. The statistical nature of these distributions has
been a concern of macroecology (Brown 1995; Gaston and
Blackburn 2000; Gaston 2003), but factors that constrain
distributions within regions are poorly understood. Or-
dinations of species across sample locations often have
high dimensionality, indicating multiple influences and
possibly idiosyncrasy and randomness in distributions;
correlations of ordinated positions of species with envi-
ronmental variables tend to account for little of the total
variance in the species distributions (e.g., Eilu et al. 2004;
Svenning et al. 2004).
The characteristic dimension (vi) of the distribution of
species i within a region is influenced by the range of
ecological conditions to which individuals are adapted
(Brown 1984; Gaston 2003); genetic differentiation of local
populations within the species’ range (Thompson 2005);
accessibility of ecologically suitable areas relative to dis-
persal capabilities of a species (Pulliam 2000; Svenning
and Skov 2004); historical changes in the distribution of
ecological zones within a region (Brown et al. 2000; Linder
2005; Galley and Linder 2006); ancestral connections with
related species within a region (Barraclough and Vogler
2000; Wiens and Graham 2005; Lovette and Hochachka
2006; Wiens et al. 2007); and interactions with competi-
tors, predators, and pathogens, as well as species in mu-
tually beneficial associations (Belyea and Lancaster 1999;
Kelt and Brown 1999; Weiher and Keddy 1999; Brown et
al. 2000). Patterns of diversity within regions integrate
mechanisms that produce new species, cause extinction,
and effect the ecological and geographic sorting of species.
At the beginning of this essay, I asserted that the dis-
tribution of species within a region is more fundamental
biologically than the coexistence of many species at a point.
This assertion implies that, in focusing on local diversity,
we have been asking the wrong questions or, perhaps, the
right questions on the wrong scale. New understanding in
community ecology will come from investigating factors
that influence the distributions of species across space and
across gradients of ecological conditions. The task is made
difficult by the fact that each species may evolve a unique
relationship to the environment, leading to idiosyncratic
distributions that bear no general relationship to environ-
mental conditions. Moreover, species interact within the
regional as well as the local context, and the packing of
species into ecogeographic space bears an unknown re-
lationship to their packing into local ecological niche
space. Indeed, the partitioning of local space might be
considered a by-product of these larger-scale processes.
Species distributions reflect the interaction of the ad-
aptations of individual species in the ecogeographic setting
of a region, including historical changes in the distribution
of environments and dispersal barriers and corridors. The
region itself can be described in terms of the range and
geographic distribution of its environments. The relation-
ships of species to the environment can be as varied as
the ways in which species can become specialized. None-
theless, phylogenetic conservatism of habitat requirements
and distribution patterns can organize the bewildering va-
riety of patterns and provide clues to dominant influences
of species characteristics on distributions. The influence
of species interactions, especially competition, on distri-
butions might be sorted out, in part, through the rela-
tionship of distributional overlap to niche separation. Most
of these approaches are incorporated into the varied re-
search programs of contemporary ecology but have yet to
be integrated into a restructuring of community ecology.
What about the use of the word “community”? It should
be kept, because populations do interact within entire
regions, but it should be associated with an expanded con-
cept of the historical and spatial dimensions of these in-
teractions. We should acknowledge that populations are
the primary entities in community ecology and that the
region is the appropriate scale for an ecological and evo-
lutionary concept of community.
Acknowledgments
I am grateful to the American Society of Naturalists for
bestowing the Sewall Wright Award and for providing an
opportunity to write this essay. J. Chase, H. V. Cornell, S.
Harrison, J. B. Losos, M. McPeek, S. Normand, S. S. Ren-
ner, K. Roy, and U. Treier provided many helpful com-
ments and suggestions on the manuscript. I thank the
College of Arts and Sciences, University of Missouri–St.

Louis, for providing a research leave, during which this
manuscript was written.
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